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SITE-SPECIFIC AND LANDSCAPE FEATURES ASSOCIATED
WITH SHRUBLAND BIRD OCCURENCE IN ANTHROPOGENIC
SHRUBLANDS IN THE NORTHEASTERN UNITED STATES
BY
RANDY SHOE
Wildlife Conservation and Biology BS, University of New Hampshire, 2015
THESIS
Submitted to the University of New Hampshire
in Partial Fulfillment of
the Requirements for the Degree of
Master of Science
in
Natural Resources:
Wildlife and Conservation Biology
December 2018
ii
This thesis has been examined and approved in partial fulfillment of the requirements for the
degree of Master of Science in Natural Resources: Wildlife and Conservation Biology by:
Thesis Director, John Litvaitis, Ph.D., Professor Emeritus
Thomas Lee, Ph.D., Associate Professor, Natural Resources & The Environment
Matthew Tarr, Ph.D., Extension Professor, UNH Cooperative Extension
Mariko Yamasaki, U.S. Forest Service, Research Wildlife Biologist
Don Keirstead, Natural Resources Conservation Service, State Resource Conservationist
On 29 November 2018
Original approval signatures are on file with the University of New Hampshire Graduate School.
iii
ACKNOWLEDGEMENTS
I would like to thank the Nuttall Ornithological Club, the Natural Resources Conservation
Service, the New Hampshire Fish and Game Department, and the University of New Hampshire
for their generous financial support. I would like to acknowledge my committee and the
following people who made this possible: Jack and Kamron Hernandez, Erin Smith, Kyle Crafts,
Jennifer Gibson, Ryan Brown, Elizabeth Godin, Mike Doherty, Casey Coupe, Cooper Bryer,
Brittany Taylor, Rory Carroll, Avery Shoe-Ricker, Amir Kirata, Beau Garcia, and Shane Bradt.
Some special thanks go to my right-hand support and friend, Erica Holm. I would like to express
my thanks and love to my daughters, Stacy and Stefanie, and my wife Shirley for their support
and love during this project; for my success would be nothing without them.
iv
LIST OF FIGURES
FIGURE 1. Distribution of 101 anthropogenic shrublands surveyed for shrubland birds in
southeastern New Hampshire, 2015-2016.
FIGURE 2. Comparing the likelihood that each of eight focal shrubland bird species was
detected within each of four anthropogenic shrubland types occurring in Rockingham and
Strafford Counties, New Hampshire May-August 2015 and 2016.
FIGURE 3. Best NPMR models predicting focal bird occurrence and bird sensitivity to an
increasing proportion of site-specific and surrounding landscape features.
FIGURE 4. The proportion of the site-specific features (SSF) and surrounding landscape
features (SLF) comprising the best fit NPMR models predicting focal species occurrence in 101
anthropogenic shrublands in southeastern New Hampshire 2015-2016.
FIGURE 5. Average percent change in predicted occurrence for eight focal shrubland bird
species in response to increasing proportions of Shrubland, Field/Pasture, and Urban
Development in the surrounding landscape (50 m – 10 km) from the best fit NPMR models.
FIGURE 6. Nonmetric multidimensional scaling (NMDS) of eight focal shrubland bird species
in a community structure of 101 anthropogenic shrublands in southeastern New Hampshire,
USA, 2015-2016.
FIGURE 7. NPMR comparing predicted occurrence of eight focal shrubland bird species in
response to increasing Vegetation Density estimated in 101 anthropogenic shrublands in
Strafford and Rockingham Counties New Hampshire 2015-2016.
FIGURE 8. NPMR comparing predicted occurrence of eight focal shrubland bird species in
response to increasing Shrubland Size (ha) estimated in 101 anthropogenic shrublands in
Strafford and Rockingham Counties New Hampshire 2015-2016.
TABLE 1. LANDFIRE 2014 Existing Vegetation Height (30 m pixels) categories grouped into
seven surrounding landscape features with reasons for grouping.
v
LIST OF TABLES
TABLE 1. LANDFIRE 2014 Existing Vegetation Height (30 m pixels) categories grouped into
seven surrounding landscape features with reasons for grouping.
vi
ABSTRACT
SITE-SPECIFIC AND LANDSCAPE FEATURES ASSOCIATED WITH SHRUBLAND BIRD
OCCURRENCE IN ANTHROPOGENIC SHRUBLANDS IN THE NORTHEASTERN
UNITED STATES
By
Randy Shoe
University of New Hampshire, December 2018
Habitats dominated by low-growing trees and shrubs are becoming increasingly uncommon in
the northeastern U.S. Human development, altered natural-disturbance regimes, and forest
succession have reduced the quantity and quality of these shrublands. As a result, over half of the
shrubland-dependent songbirds in the region have experienced long-term population declines.
Anthropogenic shrublands, including regenerating clearcuts, sand and gravel mines, old fields,
and transmission line rights-of-way may provide nesting habitat for most shrubland birds; but
differences in size, site-specific features, and landscape composition may affect bird use. To
assess the features that may influence shrubland bird occurrence in anthropogenic shrublands, I
conducted presence/absence surveys of 8 species [alder flycatcher (Empidonax alnorum), brown
thrasher (Toxostoma rufum), blue-winged warbler (Vermivora cyanoptera), chestnut-sided
warbler (Setophaga pensylvanica), eastern towhee (Pipilo erythrophthalmus), field sparrow
(Spizella pusilla), indigo bunting (Passerina cyanea), and prairie warbler (Setophaga discolor)]
in 101 sites in southeastern New Hampshire during the 2015 and 2016 nesting seasons. For each
shrubland, I measured area, site-specific features (e.g., vegetation height, density, and coverage),
and characteristics of surrounding landscape features within different buffer zones. Overall, 67%
of the variables in the best models predicting bird occurrence were landscape features and 33%
were site-specific features. Bird occurrence at a site was positively associated with the proportion
vii
of shrublands in the surrounding landscape, particularly within a 500 m buffer. Occurrence of all
species except blue-winged warblers and indigo buntings was negatively associated with the
proportion of urban development in the surrounding landscape. Shrubland bird species richness
increased with vegetation density until vegetation density became too dense for brown thrashers,
field sparrows, and prairie warblers. Occurrence of all species except blue-winged warblers
increased with shrubland size. These results provide opportunities to enhance existing
anthropogenic habitats to benefit populations of declining shrubland birds.
viii
TABLE OF CONTENTS
ACKNOWLEDGEMENTS………………………………………………….. iii
LIST OF FIGURES………………….……….………………………………. iv
LIST OF TABLES…………………………………………………...……….. v
ABSTRACT………………………………………………………………….. vi
CHAPTER PAGE
1. INTRODUCTION…………………………………………………….…….. 1
2. METHODS…………………………………………………………….……. 6
2.1. Study Area and Shrubland Selection……………………..………….…. 6
2.2. Focal Species……………………………………………………….…… 8
2.3. Field Methods………………………………………………….……….. 9
2.3.1. Site-Specific Features ………………………….……….……….. 9
2.3.2. Landscape Features ………………….………….……….………. 11
2.4. Statistical Analysis……………………………………………………… 13
2.4.1. Bird Occurrence …………………………………………….…… 13
2.4.2. Site-specific and Landscape Features Affecting Occurrence …… 13
3. RESULTS…………………………………………………………………… 15
3.1. Shrubland Bird Detection and Shrubland Types …………….………… 15
3.2. Bird Responses to Site-Specific and Landscape Features …………...… 16
3.3. Bird Reponses to Shrubland Size ……………………………………… 23
4. DISCUSSION……………………………………………………………….. 24
4.1. Importance of Landscape Composition to Breeding Shrubland Birds ……. 25
4.2. Vegetation Structure, not Species Composition, Predicted Shrubland Bird
Occurrence. …………………………………………………………….. 28
4.3. Shrubland Birds Occur More Predictably in Larger Shrublands ……...…... 32
5. CONCLUSION AND MANAGEMENT RECOMMENDATIONS ………. 33
LITERATURE CITED………………………………………….……………… 34
ix
APPENDIX A. Shrubland site-specific features, ranges and averages for sand
and gravel mines (S&GM), transmission line rights-of-way
(ROW), old fields (OF), and clearcuts (CC)…………………… 41
APPENDIX B. Shrubland surrounding landscape features, ranges and averages
for sand and gravel mines (S&GM), transmission line
rights-of-way (ROW), old fields (OF), and clearcuts (CC)……. 42
APPENDIX C. UNH Institutional Animal Care and Use Committee approval
letter …………………………………………………………… 43
1
1. INTRODUCTION
Habitats dominated by low-growing trees and shrubs are becoming increasingly uncommon in
the northeastern United States (Hunter et al. 2001, Motzkin and Foster 2002) where human
developments, altered disturbance regimes, and forest succession have reduced the quantity and
quality of shrublands that are important to many wildlife species (Litvaitis 1993, Askins 2001,
Thompson and DeGraaf 2001). As a result, shrubland-dependent wildlife, including 29 of 41
shrubland-dependent birds have experienced long-term population declines (Rosenberg et al.
2016). In response to these declines, federal and state natural resource agencies, in partnership
with public and private landowners, are working to create and maintain shrublands (Oehler et al.
2006). The largest and most recent of these efforts in New England and eastern New York
involves clearcutting large (> 5 ha) blocks of second growth forest near existing anthropogenic
shrublands such as sand and gravel mines, transmission line rights-of-way (ROW), and shrubby
old fields, to restore populations of New England cottontails (Sylvilagus transitionalis) (Fuller
and Tur 2012). While this effort is focused on establishing a network of shrublands that support
cottontail breeding and dispersal, these habitats and their arrangement are expected to provide
similar benefits to declining shrubland birds.
Throughout New England, populations of many shrubland bird species, including prairie
warblers (Setophaga discolor), blue-winged warblers (Vermivora cyanoptera), chestnut-sided
warblers (Setophaga pensylvanica), brown thrashers (Toxostoma, rufum), eastern towhees
(Pipilo erythrophthalmus), field sparrows (Spizella pusilla), and indigo buntings (Passerina
cyanea) rely on anthropogenic shrublands as their primary breeding habitat wherever natural
xeric shrublands such as pine barrens are absent (Dettmers 2003, Gifford et al. 2010). Here,
shrubby old fields, shrubby ROWs, active and idle sand and gravel mines, and regenerating
2
clearcuts are the most common anthropogenic habitats these birds use for breeding (Confer and
Pascoe 2003, Schlossberg and King 2007, King et al. 2009a). Although there is overlap in the
bird species expected to use each of these shrubland types (Rodewald and Vitz 2005, Bulluck
and Buehler 2006, Schlossberg and King 2007), differences in growing conditions, vegetation
structure, and plant species composition may lead to distinct bird communities among shrublands
(Skowno and Bond 2003, Bulluck and Buehler 2006, Schlossberg and King 2007). Further, the
extent to which shrubland size and shape, and landscape composition surrounding such
shrublands interact to influence habitat selection by shrubland birds has received limited study
(Askins et al. 2007, Chandler et al. 2009a, Roberts and King 2017). Understanding how these
factors influence habitat selection by shrubland birds is critical for determining the amount of
habitat currently available for these species and for guiding habitat management and
conservation efforts aimed at maintaining functioning bird populations.
Vegetation structure can have an important influence on bird community assemblages (Karr
and Roth 1971) and shrubland birds are recognized as habitat specialists that require or prefer
specific vegetation structure or composition for nesting and feeding (Schlossberg and King 2007,
King et al. 2009b, Schlossberg et al. 2010). As a result, differences in vegetation structure and
composition between anthropogenic shrublands may result in each bird species occurring in
different abundances among different shrubland types or being completely absent from
shrublands lacking required conditions (DeGraaf and Yamasaki 2003, Schlossberg and King
2007). Composition of bird communities in these habitats should also change as vegetation
structure changes (Schlossberg 2009). For example, by 10-15 years after cutting a stand of
northern hardwoods, woody vegetation has typically grown beyond the shrub stage required by
nesting shrubland birds and management is required to return the site to a shrubby condition
3
(DeGraaf and Yamasaki 2003, Dettmers 2003). Further, many ROWs in New England are
maintained in a shrubby condition by mowing with a brontosaurus-style forestry mower (a large
flailing-head mower attached to an excavator used to grind up shrubs and young trees) every 3-4
years to cut tree saplings that will grow tall enough to intercept the transmission lines (K.
Nelson, Eversource, personal communication). Immediately following mowing, the vegetation
along a ROW is typically dense herbaceous cover interspersed with low, scattered shrubs that
quickly succeed to very tall and dense shrubs and tree saplings during the 3-4 years between
mowing events. This succession of vegetation can result in wide variation in the composition and
density of the shrubland bird community within a ROW throughout a mowing cycle (Kroodsma
1982, King and Byers 2002, Askins et al. 2012). Understanding how differences in shrubland
vegetation structure influences the occurrence of different shrubland bird species is an important
step toward determining how different anthropogenic shrublands function as habitat.
Although vegetation structure is generally considered more important than plant species
composition when birds select habitats (MacArthur et al. 1962, James 1971, Karr and Roth
1971), shrub species composition may influence habitat use and selection by shrubland birds
(Willson 1974, Rotenberry 1985, Schlossberg et al. 2010). For example, preference for nesting in
non-native invasive shrubs vs. native shrubs varies among areas (Stoleson and Finch 2001,
Heckscher 2004, Schlossberg and King 2010). Differences in food resources, particularly insects
(Burghardt et al. 2010, Pendleton et al. 2011, Fickenscher et al. 2014) and fruits (Howe and
Estabrook 1977, Herrera 1982, Smith et al. 2013) can also vary between habitats dominated by
non-native vs. native plants and contribute to differences in the bird community among habitats
(Burghardt et al. 2009, Tarr 2017, E. Holm, University of New Hampshire, unpublished data).
Understanding whether shrubland birds select for or avoid anthropogenic shrublands containing
4
non-native shrubs is important for understanding differences in bird distribution among
shrublands and for guiding habitat management decisions regarding if and when non-native
shrubs should be controlled to benefit birds.
Factors other than vegetation structure and composition may also have an important influence
on shrubland bird use of anthropogenic shrublands. Specifically, the size and shape of shrublands
may be especially important because most shrubland birds are considered to be area sensitive
(Rodewald and Vitz 2005, Lehnen and Rodewald 2009a, Shake et al. 2012) and absent from
openings below some minimum size (Taylor and Taylor 1979, Costello et al. 2000, Askins et al.
2007). For example, Askins et al. (2007) suggested a minimum shrubland size of 0.6 ha for
shrubland birds and Costello et al. (2000) found clearcuts < 0.8 ha were too small to support
most shrubland bird species in northern New Hampshire. Larger shrublands are likely to support
greater shrubland bird richness due, in part, to the greater diversity of microhabitats they contain
compared to smaller openings (Rudnicky and Hunter 1993), but shrubland openings < 1.1 ha can
provide habitat for some shrubland birds if those openings contain required microhabitats
(Roberts and King 2017). Similarly, wide ROWs tend to support a greater diversity of shrubland
birds than narrow ROWs (Anderson et al. 1977, Confer and Pascoe 2003, Askins et al. 2012).
This area sensitivity may also be the result of shrubland birds preferring to nest away from
habitat edges where predation risk can be greater than within shrubland interiors (Shake et al.
2011). As a result, long and narrow shrublands (i.e., ROWs) may support a lower diversity and
abundance of shrubland birds than large and comparatively wider openings such as clearcuts
(Anderson et al. 1977, Schlossberg and King 2007, Askins et al. 2012), so differences in
shrubland size and shape may influence shrubland bird occurrence among anthropogenic
shrubland types.
5
Finally, bird habitat selection may also be influenced by the composition of the landscape
surrounding a site (Hinsley et al. 1995, Askins et al. 2012). Few studies have investigated the
importance of landscape composition to predict shrubland bird occurrence (e.g., Askins et al.
2007, Chandler et al. 2009b, Shake et al. 2012) and the results of these studies seem equivocal.
For example, in landscapes composed predominately by forest, wetlands, or shrublands (i.e., not
human-developed landscapes), Hagan and Meehan (2002) found that landscape composition
within 1 km of shrublands explained the presence/absence of only 2 out of 20 bird species and
Chandler (2006) found shrubland bird occurrences in wildlife openings was better predicted by
microhabitat variables within openings than by the proportion of shrublands in the surrounding
landscape. These results may be due, in part, to shrubland birds evolving stronger associations to
specific vegetation conditions within shrublands rather than to the composition of the matrix
landscape that was historically composed predominantly of mature forest (Askins et al. 2007).
However, in predominately developed landscapes, the dominant landcover may be especially
important for predicting shrubland bird occurrence in remnant shrublands (Roberts and King
2017), with the importance of the surrounding landscape composition extending beyond 100 m
from study shrublands (Hostetler and Knowles-Yanez 2003). As suburban development
increases in many areas of New England, understanding how landscape habitat composition
influences shrubland bird distribution among shrublands is important for planning landscape-
level habitat management and conservation to benefit declining bird species.
To examine the role of site-specific features (e.g., vegetation composition and shrubland size)
and surrounding landscape composition, I examined the occurrence of common shrubland birds
in four types of anthropogenic shrublands; clearcuts, sand and gravel mines, old fields, and
ROWs. I predicted that site-specific features would have a greater influence on shrubland bird
6
occurrences than surrounding landscape features, that surrounding landscape features within 500
m of a shrubland would have a greater influence on bird occurrences than features farther away,
and that bird occurrences would be associated positively with the proportion of shrubland and
negatively with the proportion of urban development within the surrounding landscape.
2. METHODS
2.1. Study Area and Shrubland Selection
I conducted this study in seacoast of southeastern New Hampshire, in Rockingham and
Strafford counties (Fig 1). The study area (2873 km2) contains approximately 70% Forest, 14%
Urban Development, 7% Fields/Pastures (i.e., habitats composed predominantly of herbaceous
vegetation < 1m tall), 4% Open Water, 4% Shrubland (i.e., habitats composed of thickets of
shrubs and young trees mixed with scattered grasses and wildflowers), 0.2% Tidal Vegetation
(i.e., habitats composed of grasses and herbaceous vegetation influenced by salt water influx)
and 0.2% Agricultural (i.e., lands managed for crops). The dominant forest type in this area is
eastern hemlock (Tsuga canadensis) – American beech (Fagus grandifolia) - oak (Quercus spp.)
- white pine (Pinus strobus) forest (Sperduto and Nichols 2012). There is more urbanization,
open water, and tidal vegetation near the coast leading into a more forested landscape as the
distance increases inland. Fields/Pastures and Agricultural landscape features are scattered
throughout the landscape as soil conditions permit.
The seacoast area of New Hampshire has unique anthropogenic shrublands, such as sand and
gravel mines, clearcuts, and ROW due to geological conditions (Sperduto and Nichols 2012),
New England cottontail habitat management (Fuller and Tur 2012), and urbanization
respectively. Old fields are less abundant due to forest succession (Litvaitis 1993). A total of 101
7
shrublands were selected as study sites: 26 clearcuts, 25 sand and gravel mines, 25 old fields, and
25 ROWs. Each shrubland was ≥ 0.8 ha and at least 250 m from the next nearest shrubland
surveyed. All ROWs were > 50 m wide and characterized by extensive shrub and sapling
hardwood growth that varied in density and vertical structure depending on time since it was
mowed. All ROWs were mowed with a brontosaurus-style forestry mower within the previous 1-
4 years and all vegetation was < 5 m tall. Study sites within ROWs varied from 1.1 to 26.5 ha
and each was considered a continuous shrubland until it was bisected by a road, parking lot,
housing development, agricultural field, or by a water feature that observers could not cross. All
silviculture openings (patch cuts, clearcuts, group selection cuts) were considered clearcuts and
were within 1-15 years of complete post-timber harvest and ranged from 1.1 to 23.7 ha. All
clearcuts had few standing trees, a hard forest/shrubland edge, and contained shrubs,
regenerating trees, and woody debris resulting in extensive vertical and horizontal diversity. Old
fields varied from 1.1 to 21.6 ha and were last mowed or grazed > 2 years prior to the study,
creating a variable vertical structure (0.2 to < 3 m tall) of dense grasses interspersed with shrubs
and small saplings. Sand and gravel mines varied in size from 1.7 to 90.9 ha and contained
extensive areas of overturned soils in a xeric environment, with pockets of shrubs, grasses and
saplings, and ponds of variable sizes created by mining activities. Active sand and gravel mines
(n = 11) were composed of a perimeter of shrubby habitat interspersed among areas where gravel
was being extracted. Idle sand and gravel mines (n = 14) were composed of a perimeter of taller
shrubs and small trees, and shrubby interiors interspersed with areas of bare ground.
8
FIGURE 1. Distribution of 101 anthropogenic shrublands surveyed for shrubland birds in
southeastern New Hampshire, 2015-2016.
2.2. Focal Species
Eight shrubland bird species were the focal species of this study: alder flycatchers, brown
thrashers, blue-winged warblers, chestnut-sided warblers, eastern towhees, field sparrows, indigo
buntings, and prairie warblers. These species are identified as “Stewardship Species of
Continental Importance” for the U.S. and Canada (Rosenberg et al. 2016), as “Species of
Greatest Conservation Need” in one or more New England state wildlife action plans, and they
require shrublands as their primary breeding habitat (Schlossberg and King 2007). Collectively,
these species require or prefer a range of shrubland habitat conditions (e.g., mesic, xeric, exposed
soils, dense/sparse shrub and tree cover, minimum required shrubland size) and they all have
experienced population declines in New England over the last 50 years (U.S. Fish and Wildlife
Service 2008, Rosenberg et al. 2016).
9
2.3. Field Methods
I conducted three presence/absence surveys for focal bird species in each shrubland between
May and August, with approximately half of the shrublands surveyed in 2015 and the other half
in 2016. Consecutive surveys at each shrubland were separated by ≥ 2 weeks. Each survey
consisted of a surveyor using a handheld speaker to broadcast the breeding songs of each focal
bird species throughout the entire shrubland to ensure complete coverage. Each focal species was
recorded as either present (detected by sight or sound) or absent (not detected) during each
survey. Once a species was detected, no further breeding song was broadcast for that species
during that survey. A species occurred at a site if it was present during at least one survey.
Surveys were conducted in the morning between sunrise and 1200 and they were not conducted
during rain, excessive wind (> 4 in Beaufort scale), fog (visibility < 200 m), or excessive heat
index (> 32 C).
2.3.1. Site-Specific Features
Eleven site-specific features were used to describe structural characteristics of sites
inventoried for breeding birds (Appendix A). Size and perimeter were determined for clearcuts,
sand and gravel mines, and old fields by walking the perimeter of the area with a handheld GPS
unit and calculating area in ArcGIS 10.2.2. I estimated size for ROWs from digital aerial
photographs in ArcGIS 10.2.2. Bare Ground Cover (the litter layer, rocks, and tree stumps), area
of Open Water, and the height, and percent cover of six vegetation classes (grasses, ferns, forbs,
native shrubs, non-native shrubs and trees) were estimated at 30 sampling points located
randomly within each shrubland. A mil-acre plot was established at each point using a 1.13 m
long rope attached to a pole held vertically at the center of the point, with the pole marked at 0.2
m intervals. The height of each vegetation class was estimated as the tallest specimen
10
representing each class (nearest 0.2 m, continuously from 0 to > 3 m). Any class missing or < 0.2
m tall was recorded as zero. Woody plants < 7.0 cm in diameter at breast height were considered
shrubs and those larger as trees. Visual estimates of the percent cover were made for bare
ground, water, and each vegetation class by looking directly down upon the plot. Cover estimates
were averaged from estimates collected simultaneously by ≥ 2 observers trained at the beginning
of each study season and weekly thereafter, to ensure consistent estimates among observers.
Because vegetation height correlated with percent cover for each vegetation class (r > 0.7), it was
not included as a model variable. Vegetation Density (all vegetation classes combined) in each
plot was measured with a density comparison gauge (5.1 cm x 7.6 cm x 10.2 cm down spout
adapter with 9 equidistant sections created by wire on the rectangular section). The density
comparison gauge was held directly against the eye 1 m above the ground and any portion of a
section containing vegetation within the 1 mil acre plot was recorded. Using the density
comparison gauge, scores for density were recorded as follows: 0 = no section with vegetation, 1
= 1 – 3 vegetated sections, 2 = 4 – 6 vegetated sections and 3 = 7 – 9 vegetated sections. Density
readings were taken at cardinal headings and averaged for each plot. Vegetation Density is based
on a combination of average percent cover or all vegetation types and the average height of
vegetation at 1 meter and categorized as: Sparse Vegetation ( ≥ 60% bare ground with limited
shrub, grass, and forb cover that seldom exceed 1 m in height); Dense Short Vegetation (60 -
80% shrub cover interspersed with grasses, forbs, and shrubs all occasionally exceeding 1 meter
in height with ≤ 40% bare ground and some small saplings < 3 m tall); Very Dense Vegetation (>
80% shrub cover with most grasses, forbs, shrubs, and saplings exceeding 1 meter in height, with
≤ 20% bare ground and some small trees > 3m tall); and All Vegetation Very Dense & Tall (>
11
80% shrub cover with all vegetation types averaging > 1m tall, some trees > 3m tall , and
minimal to no bare ground).
2.3.2. Landscape Features
To estimate the landscape composition surrounding each shrubland, six buffer distances (50
m, 250 m, 500 m, 1 km, 5 km, and 10 km) were establish around each shrubland using ArcGIS
10.2.2. The 28 LANDFIRE 2014 Existing Vegetation Height (30 m pixels) (LANDFIRE 2014)
data land categories were combined into seven surrounding landscape features and then
estimated within each buffer distance (Appendix B). Specifically, I estimated the proportion of
the landscape that was Agriculture, Field/Pasture, Open Water, Shrubland, Tidal Vegetation,
Forest, and Urban Development within each buffer. Although the larger buffer landscape
features incorporated all subsequent smaller buffer landscape features, all surrounding landscape
features within each buffer were retained in the analyses regardless of correlation strength due to
buffer overlap. Variability in the percentages of surrounding landscape features decreases as
buffer distances decrease around the shrublands.
I was unable to examine all vegetation types in every buffer distance and categorizing
LANDFIRE data into surrounding landscape features had its challenges. Growth of vegetation
over time, misinterpreted vegetation types within LANDFIRE, and the combining of
LANDFIRE into only seven types of surrounding landscape features may affect the responses of
the shrubland birds. I found most representations of LANDFIRE data did match the vegetation at
the surveyed sites after two years or was within the appropriate categories I selected as
surrounding landscape features.
12
TABLE 1. LANDFIRE 2014 Existing Vegetation Height (30 m pixels) categories grouped into
seven surrounding landscape features with reasons for grouping.
LANDFIRE 2014 category Surrounding landscape
feature category
Reason
Barren Shrubland Sparse shrub vegetation
Developed-Roads Urban Development Urban development
Developed-Upland Deciduous Forest Trees Trees in developed areas
Developed-Upland Evergreen Forest Trees Trees in rural and developed areas
Developed-Upland Herbaceous Fields, Pastures Golf courses/fields: mowed often
Developed-Upland Mixed Forest Trees Tree pockets in developed areas
Developed-Upland Shrubland Urban Development Small backyards by rural roads
Developed-High Intensity Urban Development Urban development
Developed-Medium Intensity Urban Development Urban development
Developed-Low Intensity Urban Development Urban development
Forest Height 0 to 5 meters Forest Forest
Forest Height 5 to 10 meters Forest Forest
Forest Height 10 to 25 meters Forest Forest
Forest Height 25 to 50 meters Forest Forest
Herb Height 0 to 0.5 meters Tidal Areas next to rivers, ponds, lakes
Herb Height 0.5 to 1.0 meters Fields, Pastures Most are fields, and pastures
Herb Height > 1.0 meters Fields, Pastures Most are fields, and pastures
NASS-Close Grown Crop Agriculture Agriculture
NASS-Row Crop Agriculture Agriculture
NASS-Row Crop-Close Grown Crop Agriculture Agriculture
NASS-Vineyard Agriculture Agriculture
NASS-Wheat Agriculture Agriculture
Open Water Open water Open water
Quarries-Strip Mines-Gravel Pits Shrubland Sparse shrub vegetation
Shrub Height 0 to 0.5 meters Shrubland Shrubland
Shrub Height 0.5 to 1.0 meters Shrubland Shrubland
Shrub Height > 3.0 meters Shrubland Shrubland
Sparse Vegetation Height Shrubland Shrubland
13
2.4. Statistical Analysis
2.4.1. Bird Occurrence
Differences in detection likelihood among shrubland types were analyzed in JMP Pro
13.0.0 (SAS Institute, Cary, NC) for each species using Kruskal-Wallis followed by Wilcoxon
tests among all pairs with a test for false discovery rate for significance.
2.4.2 Site-Specific and Landscape Features Affecting Occurrence
Shrubland bird associations between the species occurrence and local and landscape features
were analyzed using non-metric multidimensional scaling (NMDS) in PC-ORD v.6.19 (MjM
Software Gleneden Beach, OR). Ordination was performed with the Sorensen distance measure
in the Autopilot Slow and Thorough mode (McCune and Grace 2002), with 250 runs using both
real and randomized data. NMDS supports presence/absence, categorical, and continuous data
allowing for unconstrained analysis. The secondary matrix of shrubland and surrounding
landscape features was generally relativized to sum of squares (P = 1) to standardize to the norm
for Sorensen distance measure. In NMDS, goodness-of-fit is measured by a stress value used to
determine the number of dimensions that adequately represent sample units in ordination space
and to indicate how well the configuration matches the data (Kruskal 1964). The starting
configuration was optimized in previous ordinations to achieve the lowest final stress (8.813) and
both 2 and 3-dimensionalities were assessed. I selected the programs default joint plot cutoff (R2
= 0.20) representing the minimum value identifying a strong relationship between the
environmental variables and shrubland bird occurrences.
Focal species likelihood of occurrence in shrublands was assessed with nonparametric
multiplicative regression (NPMR) using Hyperniche v.2.30 (MjM Software, Gleneden Beach,
OR) because I expected birds would have a complex, non-linear or even an asymmetrical
14
response to variables. NPMR creates a multidimensional environmental space unconstrained by
linear responses and uses multiplicity of the variables to model species responses (McCune and
Mefford 2009). Specifically, it uses all the variables, and combinations of all variables, to target
the best location in multidimensional space identifying successively important variables that best
explain the likelihood of a species occurring (McCune and Mefford 2009). Shrubland types were
combined to determine focal species occurrence associated with surrounding landscape features.
Model form was set to Local Mean – Gaussian to center the probability density function on the
target point achieving full weighting for an observation with the same environment as the target
point. This allows a diminishing weight in observations with increasing distance from the target
point. All other settings for Free Search were set to defaults. I assessed model quality by using
the maximum cross-validated coefficient of determination R2 (xR2) from the best fit models
(McCune and Mefford 2009) and conducted sensitivity analyses. Sensitivity is expressed as a
proportion of the range of the response variable within each model (values are only compared to
each other within the model) and higher values have more influence and indicate the species is
more sensitive to the model (McCune and Mefford 2009). The minimum average neighborhood
size for acceptable model, and minimum neighborhood size for estimate, were set at automatic
and the response curves overfitting controls were set at medium. In the predicted occurrence
percent change, I considered a response to be negligible if the response achieved < 20% change
over the range of the variable.
To determine whether a species responded more to site-specific or surrounding landscape
features I totaled the number of features comprising each species’ best NPMR model and
calculated a percentage of the total composed by each category of features to determine which
was greater.
15
3. RESULTS
3.1. Shrubland Bird Detection and Shrubland Types
All but one of the eight focal species were detected at least once in every shrubland type;
brown thrashers were detected in all shrubland types except clearcuts (Fig 2). P-values are
derived from Kruskal-Wallis followed by Wilcoxon tests among all pairs with a test for false
discovery rate for significance. Brown thrashers were more likely (P < 0.008) to be detected in
sand and gravel mines than in the other shrubland types. Prairie warblers and field sparrows were
more likely (P < 0.009 for both) to be detected in ROWs and sand and gravel mines than in
clearcuts or old fields. Eastern towhee detections were greatest in ROWs and sand and gravel
mines, and the proportion of detections in ROWs was greater (P < 0.004) than that in clearcuts
and old fields. Chestnut-sided warbler detections were greatest in ROWs and clearcuts, and the
proportion of detections in ROWs was greater (P < 0.006) than those in old fields and sand and
gravel mines. Indigo bunting detections were greatest in sand and gravel mines and ROWs; the
proportion of indigo bunting detections in sand and gravel mines was greater (P < 0.002) than
those in clearcuts and those in old fields (P < 0.007); those in ROWs were greater (P < 0.019)
than in clearcuts. There was no difference in detections among shrubland types for alder
flycatchers or blue-winged warblers.
16
FIGURE 2. Comparing the likelihood that each of eight focal shrubland bird species was
detected within each of four anthropogenic shrubland types occurring in Rockingham and
Strafford Counties, New Hampshire May-August 2015 and 2016. Differences in detections
analyzed using Kruskal-Wallis followed by Wilcoxon tests among pairs with a false discovery
rate for significance. Bars with different letters differ (P < 0.05).
3.2. Bird Responses to Site-Specific and Landscape Features
Site-specific features for shrublands combined had averages for Size (9.7 ha), Perimeter (1.9
km), Vegetation Density (1.5), Species Richness (4.8), Open Water cover (0.2%), Bare Ground
cover (35.6%), Grass cover (34%), Fern cover (9.4%), Native Shrub cover (33.3%), Non-native
Shrub cover (7%), Tree cover (7.4%) and Forb cover (33.1%). All shrubland site-specific
features with ranges and averages for sand and gravel mines (S&GM), transmission line rights-
of-way (ROW), old fields (OF), and clearcuts (CC) are in Appendix A.
Overall, Urban development and Field/Pasture increase in the surrounding landscape out to
500 meters and then decrease from 1 km to 10 km. Surrounding Shrubland in the landscape
decreases as distance increases from the shrubland and there is less than a 3% change of Forest
17
in the surrounding landscape at all buffer distances (Appendix B). All shrubland surrounding
landscape feature ranges and averages for sand and gravel mines (S&GM), transmission line
rights-of-way (ROW), old fields (OF), and clearcuts (CC) are in Appendix B.
The best-fit NPMR models included site-specific and landscape features (Fig 3). Three birds
(alder flycatcher, brown thrasher, chestnut-sided warbler) were positively associated with
surrounding Shrubland. Five birds (brown thrasher, prairie warbler, eastern towhee, alder
flycatcher, field sparrow) were negatively associated with surrounding Field/Pasture and the
blue-winged warbler was positively associated with Field/Pasture. Two birds (chestnut-sided
warbler, indigo bunting) were negatively associated with surrounding Urban Development. Two
birds (field sparrow, blue-winged warbler) were positively associated with Tidal Vegetation and
two birds (indigo bunting, prairie warbler) were negatively associated with Tidal Vegetation.
Two birds (brown thrasher, field sparrow) were negatively associated with Open Water and the
prairie warbler was positively associated with Open Water. The alder flycatcher was positively
associated with Agriculture and the blue-winged warbler was negatively associated with
Agriculture. Two birds (indigo bunting, chestnut-sided warbler) were positively associated with
Vegetation Density and the brown thrasher was negatively associated with Vegetation Density.
Two birds (eastern towhee, field sparrow) were associated positively with Shrubland Size. The
blue-winged warbler was positively associated with Non-native shrubs. The alder flycatcher was
negatively associated with Bare Ground Cover and the eastern towhee was positively associated
with Bare Ground Cover. The chestnut-sided warbler was negatively associated with Tree
Cover. The prairie warbler was negatively associated with Grass Cover.
18
FIGURE 3. Best NPMR models predicting focal bird occurrence and bird sensitivity to an
increasing proportion of site-specific and surrounding landscape features. Positive (+) and
negative (-) signs indicate the direction of association of the species to the variable. Higher
sensitivity values indicate the predicted occurrence of the species changes readily
(positive/negative) in response to changes in the variable.
19
The importance of site-specific and surrounding landscape features as predictors of focal bird
occurrence differed among the species (Fig 4). Specifically, surrounding landscape features
comprised 75% of the feature types associated with alder flycatchers, brown thrashers, blue-
winged warblers, field sparrows, and prairie warblers and 67% of the feature types associated
with indigo buntings. Comparatively, the site-specific features comprised 67% of the feature
types associated with eastern towhees. The site-specific and surrounding landscape features each
comprised 50% of the feature types associated with chestnut-sided warblers (Fig 4). These
results were derived from the best models that explained the occurrences of each focal species
(Fig 3).
FIGURE 4. The proportion of the site-specific features (SSF) and surrounding landscape
features (SLF) comprising the best fit NPMR models predicting focal species occurrence in 101
anthropogenic shrublands in southeastern New Hampshire 2015-2016.
Combined, Shrubland, Field/Pasture, and Urban Development account for 55% of
surrounding landscape features identified in the best fit models predicting the occurrence for all
species. The average percent change in predicted occurrence of all species was associated
positively with increasing Shrubland and variable, by species, with increasing Field/Pasture and
20
Urban Development across all buffer distances (Fig 5). Specifically, alder flycatchers, blue-
winged warblers and chestnut-sided warblers responded positively, and all other species
responded negatively, to Field/Pasture. Blue-winged warblers and indigo buntings responded
positively, and all other species responded negatively, to Urban Development.
FIGURE 5. Average percent change in predicted occurrence for eight focal shrubland bird
species in response to increasing proportions of Shrubland, Field/Pasture, and Urban
Development in the surrounding landscape (50 m – 10 km) from the best fit NPMR models.
Positive value indicates an increase in average predicted occurrence and negative value indicates
a decrease in average predicted occurrence.
A convergent three-dimensional solution was reached by NMDS comparing site-specific and
surrounding landscape features to shrubland bird occurrence (Fig 6). The three axes captured a
total of 91.8% of the variance in the model (axis 1: 44.8%, axis 2: 33.4%, axis 3: 13.5%). Axes 1
and 2 had the highest coefficients of determination and explained 78.2% of the shrubland bird
occurrence in relation to the site-specific and surrounding landscape feature data. Species
richness (R2 = 0.320) and Field/Pasture within 1 km (R2 = 0.211) met the minimum association
of R2 ≥ 0.2 and had the strongest relationship between the site-specific and surrounding
21
landscape features to the ordination scores in ordination space. Specifically, the chestnut-sided
warblers and blue-winged warblers had positive associations with Field/Pasture within 1 km as
it increased from axis 1, and Species richness increased as Vegetation density increased (Fig 6).
FIGURE 6. Nonmetric multidimensional scaling (NMDS) of eight focal shrubland bird species
in a community structure of 101 anthropogenic shrublands in southeastern New Hampshire,
USA, 2015-2016. Field/Pasture within 1 km and Species richness vectors are oriented toward the
direction of greatest increase and vector lengths are proportional to R2 with the ordination. The
angle between vectors indicates the correlation between variables. Convex hulls indicate
Vegetation Density categories and are defined in METHODS 2.3.1.
22
Based on the related strength of Species Richness to Vegetation Density a separate analysis
was conducted comparing Species Richness to Vegetation Density. All eight focal species were
associated with Dense Short Vegetation (n = 27) and Very Dense Vegetation (n = 38), which,
combined had the most shrublands associated with them. Species Richness in Sparse Vegetation
Density (n = 22) was limited to seven of the focal species with blue-winged warbler absent and
Species Richness in All Vegetation Very Dense and Tall (n = 14) was limited to five species with
the brown thrasher, prairie warbler and field sparrow absent.
The NPMR indicated either a positive or negative percent change in predicted occurrence to
increasing Vegetation Density (Fig 7). The alder flycatchers, blue-winged warblers, indigo
buntings, chestnut-sided warblers and eastern towhees had an overall positive predicted
occurrence as Vegetation Density increased. The brown thrashers, field sparrows, and prairie
warblers had and overall negative predicted occurrence as Vegetation Density increased.
23
FIGURE 7. NPMR comparing predicted occurrence of eight focal shrubland bird species in
response to increasing Vegetation Density estimated in 101 anthropogenic shrublands in
Strafford and Rockingham Counties New Hampshire 2015-2016. Percent increase/decrease in
predicted occurrence is indicated in parentheses. Alder flycatchers, brown thrashers, blue-winged
warblers, chestnut-sided warblers, indigo buntings, and prairie warblers have significant (>20%
change) predicted occurrence responses.
3.3. Bird Responses to Shrubland Size
The NPMR model indicated either a positive or negative percent change (% change) in
predicted occurrence of the eight focal shrubland bird species in response to increasing
Shrubland Size and the association was generally positive (Fig 8). For alder flycatchers (%
change = 43.0), brown thrashers (% change = 44.2), eastern towhees (% change = 31.1), field
sparrows (% change = 53.1), prairie warblers (% change = 27.4), and indigo buntings (% change
= 9.2) the predicted percent change in response to increasing Shrubland Size was clearly and
consistently positive. Blue-winged warblers (% change = -4.3) had a minimal negative response
to increasing Shrubland Size. Chestnut-sided warblers (% change = 1.1) had a positive response
to increasing Shrubland Size that peaked around 7.8 ha in Shrubland Size, then the response was
negative as Shrubland Size increased. In the NPMR model (Fig 8), the response lines for the
indigo buntings and brown thrashers are longer because a greater number of predicted
24
occurrence responses in sand and gravel mines met the minimum neighborhood size for
inclusion in the analysis.
FIGURE 8. NPMR comparing predicted occurrence of eight focal shrubland bird species in
response to increasing Shrubland Size (ha) estimated in 101 anthropogenic shrublands in
Strafford and Rockingham Counties New Hampshire 2015-2016. Percent increase/decrease in
predicted occurrence is indicated in parentheses.
4. DISCUSSION
Contrary to previous studies (Bulluck and Buehler 2006, Chandler 2006, Shake et al. 2012), I
found that shrubland bird occurrence was better predicted by the composition of the surrounding
landscape than by habitat features measured within shrublands. But similar to those and other
studies (King et al. 2009b, Bauer 2018), shrubland bird occurrence and species richness were
associated positively with vegetation density within shrublands and shrubland size. Specifically,
shrubland birds as a group were more commonly present in shrublands situated in landscapes
where the surrounding 500 m was composed of a greater proportion of shrublands than of fields
or urban development, in shrublands > 12 ha, and in shrublands with Dense Short Vegetation or
25
Very Dense Vegetation. These results can serve as a basic guide for identifying, managing and
conserving shrublands that are likely to support the greatest diversity of shrubland birds within
contemporary landscapes. However, species-specific preferences for specific microhabitats (e.g.,
sparse/dense/short/tall vegetation, amount of bare ground, mesic/xeric soils) indicates each bird
species will occur most predictability in shrublands that support its preferred microhabitats, and
local landscapes composed of multiple shrubland types (e.g., ROW, clearcuts, old fields, sand
and gravel mines, pine barrens, beaver meadows) occurring across a range of growing conditions
(e.g., mesic, xeric), should support the greatest diversity and abundance of shrubland birds
(Chandler et al. 2009b, King et al. 2009a, Gifford et al. 2010). Overall, sand and gravel mines
and ROW were the largest shrublands in my study and they supported both the greatest diversity
of microhabitats and the greatest species richness of shrubland birds (Fig 2).
4.1. Importance of Landscape Composition to Breeding Shrubland Birds
Occurrence of all species was associated positively with increasing proportions of Shrublands
in the surrounding landscape. All species except the blue-winged warbler, chestnut-sided
warbler, and alder flycatcher were associated negatively with increasing proportions of
Field/Pasture and the indigo bunting and blue-winged warbler were associated positively to
increasing proportions of Urban Development in the surrounding landscape (Fig 5). The positive
response I observed for blue-winged warblers to Urban Development and Fields/Pasture (Fig 5)
is likely because these land uses were more common within 500 m around old fields, where blue-
winged warbler occurrence was greatest (Fig 2), than around the other shrubland types in my
study area. The negative response and sensitively of most focal birds to greater proportions of
Field/Pasture and Urban Development may be associated with attributes of these habitats that
can have negative consequences to breeding bird fitness. For example, fragmentation by human
26
development and agriculture can decrease nesting success due to increased nest predation
(Rudnicky and Hunter 1993, Rodewald and Yahner 2001) and/or by parasitism by brown-headed
cowbirds (Burhans and Thompson III 2006, Askins et al. 2012). Urban Development may further
reduce nesting success in shrublands due to elevated noise and light levels (Oneal and
Rotenberry 2009) and/or reduced food and cover resources compared to when the adjacent
habitat is mature forest (King and DeGraaf 2004, Oneal and Rotenberry 2009, Shake et al. 2012).
Shrubland bird fitness should be greatest in landscapes where multiple shrublands are
clustered in close proximity within a matrix composed primarily of forest and wetlands and a
low proportion of fields, pasture, agricultural crops or urban development (Roberts and King
2017). Specifically, shrubland birds are known to increase their fitness by readily moving among
adjacent habitat patches during a breeding season to seek extra-pair copulations (Nolan 1978,
Byers et al. 2004, Akresh et al. 2015), to relocate following nest failure (Best 1977, Nolan 1978),
and possibly to scout and assess other potential habitats (Nolan 1978, Whitaker and Warkentin
2010). Also, this large-scale habitat selection may also indicate shrubland birds may have more
movement during the breeding season (Lehnen and Rodewald 2009b) than previously examined.
However, the ideal distance shrublands must be from one another to facilitate these bird
movements requires further study. Roberts and King (2017) determined that small (0.02 -1.3 ha)
shrublands were more likely to be occupied by shrubland birds when those shrublands occurred
within < 100 m of larger shrublands, and Lehnen and Rodewald (2009a) recommended
clustering shrublands within 1 km of one another to facilitate bird movements.
During my study, I observed individual color-banded prairie warblers, field sparrows, and
eastern towhees movements averaging 6.7 km, and as much as 29 km, within multiple
shrublands during the same nesting season (R. Shoe, University of New Hampshire, unpublished
27
data). Although these birds used multiple shrublands juxtaposed across a large local landscape, I
found shrubland bird occurrence was best predicted by the proportion of shrublands within 500
m of my study sites, likely because the proportion of shrublands in the landscape around my sites
decreased as distance from sites increased. It is possible that shrubland bird occurrence within
500 m of my sites may be due to neighborhood fidelity, whereas a bird has fidelity to multiple
sites within a radius that supports its habitat preference. More study is needed on the possible
neighborhood fidelity of shrubland birds. My results suggest clustering shrublands within 500 m
of one another may be best for encouraging shrubland bird occurrence in anthropogenic
shrublands within a suburban landscape and may possibly support neighborhood fidelity.
Unlike the other focal species, occurrence of chestnut-sided warblers was predicted equally
by site-specific and surrounding landscape features (Fig 4). The chestnut-sided warbler is
strongly influenced by microhabitat and landscape-level variables (Hagan and Meehan 2002,
Chandler 2006) and its positive response to Vegetation Density and negative response to Tree
Cover (Fig 3) are consistent with other studies (Askins et al. 2007, 2012). Movement by
chestnut-sided warblers during the breeding season can be limited by Urban Development (Byers
et al. 2004), and this may explain the negative predicted occurrence I observed for chestnut-sided
warblers in response to Urban Development within 50 m (Fig 3). An explanation of why the
chestnut-sided warbler had a positive response to a decreasing proportion of Shrublands out to
10 km (Fig 3) may be as the amount of surrounding Shrublands decrease from the occupied
habitat and the amount of Forest increases, it may be chestnut-sided warblers are responding to
its preference of an increasing proportion of understory stems < 1m tall (Hagan and Meehan
2002), provided that the forest has limited canopy cover and sufficient understory. Similar to my
other focal species, shrublands located in a landscape with minimum Urban Development are
28
those most likely to benefit chestnut-sided warblers. Managers can further attract chestnut-sided
warblers to shrublands by encouraging microhabitats composed of dense to very dense
vegetation surrounded by forest with an understory growth.
The eastern towhee was the only focal species for which site-specific features better
predicted occurrence than the surrounding landscape features (Fig 4) and towhees were most
sensitive (positively) to Bare Ground Cover and Shrubland Size (Fig 3). Increased sensitivity to
Bare Ground Cover may be explained by the habit of eastern towhees to forage and nest on the
ground in habitats with leaf litter and dense grasses, forbs and ferns. Towhees are reported to use
shrublands that span a range of sizes (0.02 - 21 ha: Askins et al. 2007, Roberts and King 2017),
and in my study area, they seem to use shrubland openings of any size as long as the shrubland
supports their foraging and nesting habits. Towhee sensitivity to Shrubland size is likely because
they occurred most predictably in large ROW and sand and gravel mines; these large openings
supported a variety of microhabitat conditions, including areas of Bare Ground Cover, Dense
Short Shrubs, and Very Dense Shrubs that satisfy multiple towhee habitat preferences. Managers
can make shrublands more suitable for eastern towhees by ensuring a site has dense to very
dense vegetation for nesting and bare ground with leaf litter for foraging.
4.2. Vegetation Structure, not Species Composition, Predicted Shrubland Bird Occurrence
Vegetation Density within shrublands had an overall positive influence on focal bird richness
and shrublands composed of Dense Short Vegetation and Very Dense Vegetation were those
most likely to be occupied by all eight focal species. However, bird response to Vegetation
Density was species-specific. As expected, brown thrashers, prairie warblers and field sparrows
were most likely to occur in shrublands with Sparse, Dense Short, and Very Dense Vegetation
and they were least likely to occur in shrublands with All Vegetation Very Dense and Tall.
29
Comparatively, blue-winged warblers and alder flycatchers were more likely to occur in
shrublands with All Vegetation Very Dense & Tall, and they were less common in shrublands
composed of Sparse Vegetation or Dense Short Vegetation. Chestnut-sided warblers were most
predictable in shrublands composed of Dense Short Vegetation or Very Dense Vegetation and
they were least likely to occur in All Vegetation Very Dense & Tall shrublands. Indigo buntings
and eastern towhees were present in all shrublands sampled and among all species, their
occurrence seemed the least influenced by vegetation density.
Because bird responses to vegetation density was species-specific, large shrublands composed
of a variety of microhabitats that support vegetation conditions ranging from Sparse Shrub
Vegetation to All Vegetation Very Dense & Tall are those shrublands most likely to support the
greatest shrubland bird richness. In my study, sand and gravel mines and ROW’s were the largest
shrublands I surveyed, and they supported both the greatest diversity of microhabitats and the
greatest shrubland bird richness (5.4 and 5.2 respectively). Specifically, sand and gravel mines
contained microhabitats consisting of mesic and xeric areas, low-growing and tall-growing
shrubs, and extensive bare ground, and they were also the shrublands with the greatest proportion
of other shrublands in the surrounding landscape; combined, these characteristics of sand and
gravel mines likely explain why they had the highest species richness of all shrubland types
surveyed. In some landscapes, sand and gravel mines may functions as the most stable and
predictable habitats for shrubland birds due to their harsh growing conditions that naturally keep
them in a shrubby condition longer than other anthropogenic shrublands such as clearcuts and
old fields where shrubby conditions are more ephemeral (Borgegard 1990, Bulluck and Buehler
2006). Regular maintenance of ROW by utility companies creates an additional source of diverse
and predictable shrubland habitat supporting a variety of shrubland bird species (King and Byers
30
2002, King et al. 2009a, Askins et al. 2012, this study) resulting from the diverse vegetation
conditions within the length of most ROW. My study indicates sand and gravel mines and
transmission ROW should be considered important habitats that can contribute to the long-term
conservation of declining shrubland birds in suburban New England landscapes.
I suspect the type of shrubland may also affect species richness with vegetation density,
especially in sand and gravel mines and ROW. Although sand and gravel mines have the highest
species richness and four of the highest detections for the species, I believe their size and the
features in the surrounding landscape may be what affects species richness more than vegetation
density. Sand and gravel mines, with less shrubs than old fields, clearcuts and ROW, have the
most shrublands surrounding them out to 500 m more than any other shrubland type. More
shrublands surround sand and gravel mines than any other shrubland type, and the value of
shrublands is important to all the species studied (Fig 5). Active sand and gravel mines and
ROW, ranging from Sparse to Very Dense Vegetation, are the only two anthropogenic habitats I
studied maintained in a shrubby condition. Habitats maintained in a shrubby condition have
indicated over a three-year mark and recapture study (2014 – 2016) that 85% of the yearly re-
sighted or re-captured prairie warblers, field sparrow, and eastern towhees have shown site
fidelity to active sand and gravel mines and ROW (R. Shoe, University of New Hampshire,
unpublished data). ROW’s are also well connected and have varying mowing schedules and
likely offer safer or easier species movement and habitat selection of desirable vegetation
densities, thereby increasing species richness.
Managers can attempt to make each shrubland suitable for a variety of shrubland bird species
by managing for a diversity of vegetation densities that are most suited for the growing
conditions of each shrubland. It may not be possible or feasible to increase vegetation diversity
31
in a shrubland composed of homogenous growing conditions; here, the best option for increasing
vegetation diversity is to identify opportunities within the surrounding 500 m landscape where
additional shrublands can be created or maintained on sites that can support different and
complimentary vegetation conditions.
Except for blue-winged warblers, vegetation species composition did not predict the
occurrence of focal shrubland birds (Fig 3). In my study area, shrubland bird richness has been
shown to increase as the proportion of invasive shrubs (e.g., Frangula alnus, Eleagnus
umbellata, Lonicera spp.) in a shrubland increases from zero to about 55% of the total shrub
cover (Tarr 2017, Bauer 2018). My study did not indicate this response for the shrubland bird
species, likely because I measured the percent cover of native and invasive shrubs as a
proportion of total vegetation cover (including grasses, forbs, ferns, trees, bare ground, and open
water), and both Bauer (2018) and Tarr (2017) measured native/invasive shrub cover as a
proportion of total shrub cover only. As a result, my sites averaged 7% Non-native shrub cover
and 33% Native shrub cover which may explain why Species Richness increased with vegetation
structure and not vegetation species composition. Blue-winged warblers responded positively to
increasing proportions of Non-native shrub cover (Fig 3), but because non-native shrubs were
most abundant in old fields, the shrubland type where blue-winged warblers were most
predictable, I cannot conclude whether they were responding specifically to Non-native shrub
cover or the high vegetation densities of old fields, coinciding with King et al. (2009b). Overall,
my results indicate that vegetation density and height was more important for influencing
shrubland bird distribution than was the native/non-native shrub composition in shrublands.
32
4.3. Shrubland Birds Occur More Predictably in Larger Shrublands
Most bird species occurred with increasing predictability as shrubland size increased above
0.8 ha (Fig 8). I predicted bird occurrence would increase with shrubland size because larger
shrublands should provide birds with a lower risk of edge effects such as nest predation (King
and Byers 2002, DeGraaf and Yamasaki 2003), and compared to smaller shrublands, they are
more likely to support a greater variety of microhabitats that support shrubland birds (Rodewald
and Vitz 2005, Chandler et al. 2009b, Singer et al. 2012). Chestnut-sided warblers and blue-
winged warblers were the exceptions to the pattern of greater predictability in response to larger
shrubland size (Fig 8). Chestnut-sided warbler occurrence became less predictable when
shrubland size exceeded 7.8 ha (Fig 8), likely explained by the fact that most shrublands larger
than 7.8 ha in my study were sand and gravel mines where the microhabitats preferred by
chestnut sided-warblers were uncommon. Similarly, blue-winged warbler occurrence was largely
unresponsive to differences in shrubland size, likely because they occurred most predictably in
old fields (Fig 2) characterized by All Vegetation Very Dense and Tall, a microhabitat type that
was uncommon in the larger ROW and sand and gravel mines in my study. Although, by not
limiting the shrubland size during my study to a maximum value, the sand and gravel mines,
averaging 17.3 ha may have swayed the predicted occurrence of the blue-winged warbler and
chestnut-sided warbler, since the sand and gravel mines were significantly larger than all other
shrubland types except ROW’s. Managing for large shrublands has long been a conservation
strategy to increase shrubland bird species richness (DeGraaf and Yamasaki 2003, Shake et al.
2012, Roberts and King 2017) and my findings agree with this strategy.
33
5. CONCLUSION AND MANAGEMENT RECOMMENDATIONS
My study was done in the southeastern seacoast of New Hampshire and my results may not
be applicable to northern or southern areas based on forest types, soil composition and because
this area is the northernmost range of the blue-winged warblers, eastern towhees, and prairie
warblers (Dunn and Alderfer 2011). The presence and absence of the blue-winged warblers,
eastern towhees, and prairie warblers may have been affected by their availability earlier or later
in the breeding season in this northernmost range of their habitat. No shrubland bird in my study
had a modeled response to Forest in the surrounding landscape and only the chestnut-sided
warbler had a response to site-specific Tree cover (negative). I suspect in heavily forested areas
the shrubland bird occurrences would be less, as found in other studies (Costello et al. 2000,
Chandler 2006), than the shrubland bird occurrences in the seacoast of New Hampshire, possibly
based on the extent of the surrounding forest, size and type of opening, and a lack of alternate
shrublands within the surrounding landscape.
Increased shrubland size and increasing vegetation density should be considered during the
planning stages of conservation site selection for shrubland birds, as most of the shrubland birds
responded positively to an increase in Size and Vegetation Density. Because shrubland Size did
not correlate to Vegetation Density, the predicted occurrence of the shrubland birds to shrubland
Size are based on the species habitat preferences to other site-specific and surrounding landscape
features. Most species predicted occurrence were in response to surrounding landscape features
(Fig 4) yet evaluating an appropriate size for shrubland bird conservation based on surrounding
landscape features of up to 10 km is unreasonable for most applications. Focusing on the site-
specific features (Size and Vegetation Density) and adjacent surrounding landscape features
(Shrublands, Urban Development, Field/Pasture) that best predicted most species occurrences
34
may be best when making a conservation decision regarding shrubland birds. Also, the trend of
increasing Urban Development and Field/Pastures as Shrubland decreases in the adjacent
surrounding landscape needs to be examined more closely. By combining the site-specific and
adjacent surrounding landscape features the birds are responding to, managers may be able to
determine if the size of a shrubland can be ‘extended’ with limited consideration to local
surrounding landscape features to increase the shrubland.
More study is also needed for shrubland bird conservation within sand and gravel mines
and ROW’s which seem to have high value to shrubland birds. Incorporating anthropogenic
shrublands maintained in a shrubby condition, such as ROW, or slow growing for an extended
time, such as sand and gravel mines (Rehounková and Prach 2006), should be considered as
primary conservation areas for shrubland birds. Generally, increasing shrub (native or exotic)
density within the shrubland habitat increases the predicted occupancy of most of the shrubland
bird species we studied.
LITERATURE CITED
Akresh, M. E., D. I. King, and R. T. Brooks. 2015. Demographic response of a shrubland bird to
habitat creation, succession, and disturbance in a dynamic landscape. Forest Ecology and
Management 336:72–80.
Anderson, S. H., K. Mann, and H. H. Shugart. 1977. Effect of transmission-line corridors on bird
populations. American Midland Naturalist 97:216–221.
Askins, R. A. 2001. Sustaining biological diversity in early successional communities: The
challange of managing unpopular habitats. Wildlife Society Bulletin 29:407–412.
Askins, R. A., C. M. Folsom-O’Keefe, and M. C. Hardy. 2012. Effects of vegetation, corridor
width and regional land use on early successional birds on powerline corridors. PloS one
7:e31520.
<http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=3282771&tool=pmcentrez&re
ndertype=abstract>. Accessed 2 Feb 2014.
35
Askins, R. A., B. Zuckerberg, and L. Novak. 2007. Do the size and landscape context of forest
openings influence the abundance and breeding success of shrubland songbirds in southern
New England? Forest Ecology and Management 250:137–147.
Bauer, M. L. 2018. Assessing the effects of habitat restoration on shrubland specialists: Case
study on the New England cottontail and shrubland birds. Thesis, University of New
Hampshire, Durham, New Hampshire.
Best, L. B. 1977. Territory quality and mating success in the field sparrow (Spizella pusilla).
American Ornithological Society 79:192–204.
Borgegard, S. 1990. Vegetation development in abandoned gravel pits: Effects of surrounding
vegetation, substrate, and regionality. Journal of Vegetation Science 1:675–682.
Bulluck, L. P., and D. A. Buehler. 2006. Avian use of early successional habitats: Are
regenerating forests, utility right-of-ways and reclaimed surface mines the same? Forest
Ecology and Management 236:76–84.
Burghardt, K. T., D. W. Tallamy, and W. Gregory Shriver. 2009. Impact of native plants on bird
and butterfly biodiversity in suburban landscapes. Conservation Biology 23:219–224.
Burghardt, K. T., D. W. Tallamy, C. Philips, and K. J. Shropshire. 2010. Non-native plants
reduce abundance, richness, and host specialization in lepidopteran communities. Ecosphere
1:art11.
Burhans, D. E., and F. R. Thompson III. 2006. Songbird abundance and parasitism differ
between urban and rural shrublands. Ecological Society of America 16:394–405.
Byers, B. E., H. L. Mays, and D. F. Westneat. 2004. Extrapair paternity increases variability in
male reproductive success in the chestnut-sided warbler (Dendroica pensylvanica), a
socially monogamous songbird. The Auk 121:788–795.
Chandler, R. B. 2006. Early-successional shrubland bird abundance and nesting success in
managed shrublands on the White Mountain National Forest. Thesis, University of
Massachusetts. Amherst, MA.
Chandler, R. B., D. I. King, and C. C. Chandler. 2009a. Effects of management regime on the
abundance and nest survival of shrubland birds in wildlife openings in northern New
England, USA. Forest Ecology and Management 258:1669–1676.
Chandler, R. B., D. I. King, and S. Destefano. 2009b. Scrub–shrub bird habitat associations at
multiple spatial scales in beaver meadows in Massachusetts. The Auk 126:186–197.
Confer, J. L., and S. M. Pascoe. 2003. Avian communities on utility rights-of-ways and other
managed shrublands in the northeastern United States. Forest Ecology and Management
185:193–205.
36
Costello, C. A., M. Yamasaki, P. J. Pekins, W. B. Leak, and C. D. Neefus. 2000. Songbird
response to group selection harvests and clearcuts in a New Hampshire northern hardwood
forest. Forest Ecology and Management 127:41–54.
DeGraaf, R. M., and M. Yamasaki. 2003. Options for managing early-successional forest and
shrubland bird habitats in the northeastern United States. Forest Ecology and Management
185:179–191.
Dettmers, R. 2003. Status and conservation of shrubland birds in the northeastern US. Forest
Ecology and Management 185:81–93.
Dunn, J. L., and J. Alderfer. 2011. National Geographic field guide to birds of North America.
6th edition. National Geographic Society, Washington D.C.
Fickenscher, J. L., J. A. Litvaitis, T. D. Lee, and P. C. Johnson. 2014. Insect responses to
invasive shrubs: Implications to managing thicket habitats in the northeastern United States.
Forest Ecology and Management 322:127–135.
Fuller, S., and A. Tur. 2012. Conservation strategy for the New England cottontail (Sylvilagus
transitionalis).
<https://newenglandcottontail.org/sites/default/files/research_documents/conservation_strat
egy_final_12-3-12.pdf>. Accessed 13 Nov 2018.
Gifford, N. A., J. M. Deppen, and J. T. Bried. 2010. Importance of an urban pine barrens for the
conservation of early-successional shrubland birds. Landscape and Urban Planning 94:54–
62.
Hagan, J. M., and A. L. Meehan. 2002. The effectiveness of stand-level and landscape-level
variables for explaining bird occurrence in an industrial forest. Forest Science 48:231–242.
Heckscher, C. M. 2004. Veery nest sites in a Mid-Atlantic piedmont forest: Vegetative
physiognomy and use of alien shrubs. The American Midland Naturalist 151:326–337.
Herrera, C. M. 1982. Seasonal variation in the quality of fruits and diffuse coevolution between
plants and avian dispersers. Ecology 63:773–785.
Hinsley, S. A., P. E. Bellamy, I. Newton, and T. H. Sparks. 1995. Habitat and landscape factors
influencing the presence of individual breeding bird species in woodland fragments. Journal
of Avian Biology 26:94–104.
Hostetler, M., and K. Knowles-Yanez. 2003. Land use, scale, and bird distributions in the
Phoenix metropolitain area. Landscape And Urban Planning 62:55–68.
Howe, H. F., and G. F. Estabrook. 1977. On intraspecific competition for avian dispersers in
tropical trees. The American Naturalist 111:817–832.
37
Hunter, W. C., D. A. Buehler, R. A. Canterbury, J. L. Confer, and P. B. Hamel. 2001.
Conservation of disturbance-dependent birds in eastern North America. Wildlife Society
Bulletin 29:440–455.
James, F. C. 1971. Ordinations of habitat relationships among breeding birds. The Wilson
Bulletin 83:215–236.
Karr, J. R., and R. R. Roth. 1971. Vegetation structure and avian diversity in several New World
areas. The American Naturalist 105:423–435.
King, D. I., and B. E. Byers. 2002. An evaluation of powerline rights-of-way as habitat for early-
successional shrubland birds. Wildlife Society Bulletin 30:868–874.
King, D. I., R. B. Chandler, J. M. Collins, W. R. Petersen, and T. E. Lautzenheiser. 2009a.
Effects of width, edge and habitat on the abundance and nesting success of scrub-shrub
birds in powerline corridors. Biological Conservation 142:2672–2680.
King, D. I., R. B. Chandler, S. Schlossberg, and C. C. Chandler. 2009b. Habitat use and nest
success of scrub-shrub birds in wildlife and silvicultural openings in western Massachusetts,
USA. Forest Ecology and Management 257:421–426.
King, D. I., and R. M. DeGraaf. 2004. Effects of group-selection opening size on the distribution
and reproductive success of an early-successional shrubland bird. Forest Ecology and
Management 190:179–185.
Kroodsma, R. L. 1982. Bird community ecology on power-line corridors in east Tennessee.
Biological Conservation 23:79–94.
Kruskal, J. B. 1964. Nonmetric Multidimensional Scaling: A Numerical Method. Psychometrika
29:115–129.
LANDFIRE. 2014. CONUS LANDFIRE Existing vegetation height. LANDFIRE v1.4.0.
<https://www.landfire.gov/evh.php>. Accessed 7 Feb 2015.
Lehnen, S. E., and A. D. Rodewald. 2009a. Investigating area-sensitivity in shrubland birds:
Responses to patch size in a forested landscape. Forest Ecology and Management
257:2308–2316.
Lehnen, S. E., and A. D. Rodewald. 2009b. Dispersal , interpatch movements, and survival in a
shrubland breeding bird community. Journal of Field Ornithology 80:242–252.
Litvaitis, J. A. 1993. Response of early successional vertebrates to historic changes in land use.
Conservation Biology 7:866–873.
MacArthur, R. H., J. W. MacArthur, and J. Preer. 1962. On bird species diversity II. Prediction
of bird census from habitat measurements. American Naturalist 96:167–174.
38
McCune, B., and J. B. Grace. 2002. Analysis of Ecological Communities. MjM Software Design,
Gleneden Beach, Oregon, USA.
McCune, B., and M. J. Mefford. 2009. Hyperniche: Multiplicative Habitat Modeling. MjM
Software Design, Gleneden Beach, Oregon, USA.
Motzkin, G., and D. R. Foster. 2002. Grasslands, heathlands and shrublands in coastal New
England: Historical interpretations and approaches to conservation. Journal of
Biogeography 29:1569–1590.
Nolan, V. 1978. The ecology and behavior of the prairie warbler (Dendroica discolor). First
edition. The American Ornithologists Union, Lawrence, KS.
Oehler, J. D., D. F. Covell, S. Capel, and B. Long. 2006. Managing grasslands, shrublands and
young forests for wildlife. A guide for the Northeast. The Northeast Upland Habitat
Technical Committee, Westboro, MA.
Oneal, A. S., and J. T. Rotenberry. 2009. Scale-dependent habitat relations of birds in riparian
corridors in an urbanizing landscape. Landscape and Urban Planning 92:264–275.
Pendleton, R. L., B. K. Pendleton, and D. Finch. 2011. Displacement of native riparian shrubs by
woody exotics: Effects on arthropod and pollinator community composition. Natural
Resources and Environmental Issues 16:197–208.
Rehounková, K., and K. Prach. 2006. Spontaneous vegetation succession in disused gravel-sand
pits: Role of local site and landscape factors. Journal of Vegetation Science 17:583–590.
Roberts, H. P., and D. I. King. 2017. Area requirements and landscape-level factors influencing
shrubland birds. Journal of Wildlife Management 81:1298–1307.
Rodewald, A. D., and A. C. Vitz. 2005. Edge and area sensitivity of shrubland birds. The Journal
of Wildlife Management 69:681–688.
Rodewald, A. D., and R. H. Yahner. 2001. Influence of landscape composition on avian
community. Ecology 82:3493–3504.
Rosenberg, K. V., J. A. Kennedy, R. Dettmers, R. P. Ford, D. Reynolds, J. D. Alexander, C. J.
Beardmore, P. J. Blancher, R. E. Bogart, G. S. Butcher, A. F. Camfield, A. Couturier, D. W.
Demarest, W. E. Easton, J. J. Giocomo, R. H. Keller, A. E. Mini, A. O. Panjabi, D. N.
Pashley, T. D. Rich, J. M. Ruth, H. Stabins, J. Stanton, and T. Will. 2016. Partners in Flight
Landbird Conservation Plan: 2016 Revision for Canada and Continental United States.
Partners in Flight Science Committee 119 pp.
Rotenberry, J. T. 1985. The role of habitat in avian community composition: physiognomy or
floristics? Oecologia 67:213–217.
Rudnicky, T. C., and M. L. Hunter. 1993. Reversing the fragmentation perspective: Effects of
clearcut size on bird species richness in Maine. Ecological Applications 3:357–366.
39
Schlossberg, S. 2009. Site fidelity of shrubland and forest birds. The Condor 111:238–246.
Schlossberg, S., and D. I. King. 2007. Ecology and management of scrubshrub birds in New
England: A comprehensive review. USDA Natural Resources Conservation Service
Resource Inventory and Assessment Division. Amherst, MA.
Schlossberg, S., and D. I. King. 2010. Effects of invasive woody plants on avian nest site
selection and nesting success in shrublands. Animal Conservation 13:286–293.
Schlossberg, S., D. I. King, R. B. Chandler, and B. A. Mazzei. 2010. Regional synthesis of
habitat relationships in shrubland birds. Journal of Wildlife Management 74:1513–1522.
Shake, C. S., C. E. Moorman, and M. R. Burchell. 2011. Cropland edge, forest succession, and
landscape affect shrubland bird nest predation. Journal of Wildlife Management 75:825–
835.
Shake, C. S., C. E. Moorman, J. D. Riddle, and M. R. Burchell II. 2012. Influence of Patch Size
and Shape on Occupancy by Shrubland Birds. The Condor 114:268–278.
Singer, M. S., T. E. Farkas, C. M. Skorik, and K. A. Mooney. 2012. Tritrophic interactions at a
community level: Effects of host plant species quality on bird predation of caterpillars.
American Society of Naturalists 179:363–374.
Skowno, A. L., and W. J. Bond. 2003. Bird community composition in an actively managed
savanna reserve importance of vegetation structure and vegetation composition.
Biodiversity and Conservation 12:2279–2294.
Smith, S. B., S. A. DeSando, and T. Pagano. 2013. The value of native and invasive fruit-bearing
shrubs for migrating songbirds. Northeastern Naturalist 20:171–184.
Sperduto, D. D., and W. F. Nichols. 2012. Natural communities of New Hampshire. 2nd edition.
NH Natural Heritage Bureau, Concord, NH. Pub. UNH Cooperative Extension, Durham,
NH.
Stoleson, S. H., and D. M. Finch. 2001. Breeding bird use of and nesting success in exotic
Russian olive in New Mexico. The Wilson Bulletin 113:452–455.
Tarr, M. D. 2017. Effects of alien shrubs on caterpillars and shrubland-dependent passerines
within three transmission line rights-of-way in southeastern New Hampshire. Dissertation,
University of New Hampshire, Durham, NH.
Taylor, C. M., and W. E. Taylor. 1979. Birds of upland openings. Management of north central
and northeastern forests for nongame birds, workshop proceedings, 21-25 January 1979,
Minneapolis, MN. USDA Forest Service General Technical Report NC-51 Costello.
Thompson, F. R., and R. M. DeGraaf. 2001. Conservation approaches for woody, early
successional communities in the eastern United States. Wildlife Society bulletin 29:483–
494.
40
U.S. Fish and Wildlife Service. 2008. Birds of Conservation Concern 2008. United States
Department of Interior, Fish and Wildlife Service, Division of Migratory Bird Management.
Arlington, Virginia. 85 pp.
Whitaker, D. M., and I. G. Warkentin. 2010. Spatial ecology of migratory passerines on
temperate and boreal forest breeding grounds. The Auk 127:471–484.
Willson, M. F. 1974. Avian community organization and habitat structure. Ecology 55:1017–
1029.
41
APPENDIX A
Shrubland site-specific features for sand and gravel mines (S&GM), transmission line rights-of-
way (ROW), old fields (OF), and clearcuts (CC). Size range (ha) and Species Richness are
included and are not considered site-specific features.
Variable Range S&GM
Average
ROW
Average
OF
Average
CC
Average
Overall
Average
Size (ha) 1-91 17.3 7.77 6.87 6.66 9.7
Size range (ha) 2-91 1-26 1-22 1-24 9.7
Perimeter (km) 0.4-12 2.2 3.1 1.4 1.3 1.9
Vegetation density 0-3 0.82 1.78 1.79 1.54 1.5
Open Water cover (%) 0-4 0 0.3 0.3 0.2 0.2
Bare Ground cover (%) 2-85 59.6 26.6 20.9 35.3 35.6
Grass cover (%) 1-91 24.2 25.6 55.7 30.5 34
Fern cover (%) 0-48 1.2 20.2 7.2 8.8 9.4
Native Shrub cover (%) 1-82 23.8 45 27.2 37.3 33.3
Non-native shrub cover (%) 0-43 2.4 8.7 8.2 8.7 7
Tree cover (%) 0-38 7.5 4.4 5.7 11.9 7.4
Forb cover (%) 1-68 25.8 30.1 44.5 32 33.1
Species Richness 1-8 5.4 5.2 4.4 4.1 4.8
42
APPENDIX B
Shrubland surrounding landscape features, ranges and averages for sand and gravel mines
(S&GM), transmission line rights-of-way (ROW), old fields (OF), and clearcuts (CC).
Variable Range
(%)
S&GM
Averages
(%)
ROW
Averages
(%)
OF
Averages
(%)
CC
Averages
(%)
Overall
Average
(%)
50m agriculture 0-1 0.00 0.11 0.06 0.00 0.04
50m field/pasture 0-62 3.70 7.06 17.86 4.24 8.21
50m open water 0-4 0.26 0.20 0.10 0.06 0.16
50m shrubs 0-71 28.79 8.02 12.77 4.68 13.57
50m tidal vegetation 0-5 0.07 0.10 0.07 0.49 0.18
50m forest 9-100 55.82 66.46 47.08 81.63 62.75
50m urban development 0-72 11.38 18.04 22.06 8.90 15.09
250m agriculture 0-1 0.01 0.13 0.05 0.02 0.05
250m field/pasture 0-44 6.14 8.04 14.33 8.98 9.37
250m open water 0-13 0.57 1.07 0.76 0.20 0.65
250m shrubs 0-38 15.84 4.68 7.44 6.59 8.64
250m tidal vegetation 0-2 0.12 0.11 0.08 0.16 0.12
250m forest 6-99 65.21 63.17 52.72 70.29 62.85
250m urban development 0-81 12.11 22.79 24.63 13.76 18.32
500m agriculture 0-3 0.01 0.13 0.25 0.03 0.11
500m field/pasture 1-37 6.03 8.85 11.86 12.06 9.70
500m open water 0-20 0.36 1.66 1.17 0.52 0.93
500m shrubs 0-33 11.14 4.58 6.08 6.17 6.99
500m tidal vegetation 0-1 0.10 0.06 0.06 0.13 0.09
500m forest 10-92 68.12 59.78 54.78 65.04 61.93
500m urban development 0-73 14.23 24.93 25.80 16.05 20.25
1km agriculture 0-4 0.13 0.10 0.34 0.13 0.17
1km field/pasture 1-36 6.92 9.66 10.52 12.87 9.99
1km open water 0-20 0.42 2.34 1.07 1.41 1.31
1km shrubs 1-20 8.85 5.15 5.57 5.43 6.25
1km tidal vegetation 0-2 0.25 0.13 0.06 0.16 0.15
1km forest 10-91 70.29 57.19 57.01 63.61 62.03
1km urban development 3-77 13.15 25.43 25.44 16.38 20.10
5km agriculture 0 0.14 0.15 0.22 0.20 0.18
5km field/pasture 3-15 7.66 9.05 11.08 10.12 9.48
5km open water 0-18 2.22 2.90 2.33 3.47 2.73
5km shrubs 3-8 5.70 5.18 5.73 5.20 5.45
5km tidal vegetation 0-1 0.16 0.20 0.17 0.22 0.19
5km forest 32-89 71.27 61.88 58.44 62.92 63.63
5km urban development 4-42 12.85 20.64 22.03 17.87 18.35
10km agriculture 0 0.16 0.16 0.18 0.18 0.17
10km field/pasture 3-13 7.30 8.48 10.18 9.64 8.90
10km open water 1-35 3.92 3.89 4.39 5.27 4.37
10km shrubs 3-6 5.01 5.02 5.35 5.16 5.13
10km tidal vegetation 0-1 0.19 0.24 0.23 0.27 0.23
10km forest 32-85 70.57 67.81 62.27 64.58 66.30
10km urban development 5-21 12.86 14.41 17.39 14.90 14.89
43
APPENDIX C